Schaereria | |
---|---|
Schaereria fuscocinerea | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Lecanoromycetes |
Subclass: | Ostropomycetidae |
Order: | Schaereriales Lumbsch & Leavitt (2018) |
Family: | Schaereriaceae M.Choisy ex Hafellner (1984) |
Genus: | Schaereria Körb. (1855) |
Type species | |
Schaereria lugubris (A.Massal.) Körb. (1855) | |
Synonyms [1] | |
Schaereria is a genus of lichen-forming fungi. It is the sole genus in the family Schaereriaceae, which itself is the only family in the Schaereriales, an order in the subclass Ostropomycetidae of the class Lecanoromycetes. [2] Most Schaereria species are crustose lichens that live on rocks. Schaereria was first proposed by Gustav Wilhelm Körber in 1855 and was later taken up by other lichenologists despite periods of disuse.
Distinctive characteristics of species in the Schaereria genus include a crustose to squamulose thallus, ascomata (fruiting bodies) with a blackened ring and a blackish disk , and asci (spore-bearing cells) that lack tholus (a thickened part of the inner wall near the tip). The secondary chemistry of the genus produces specific substances, including the pigment known as Cinereorufa-green in Schaereria cinereorufa . The genus, having a cosmopolitan distribution, primarily favours cold to cool climates of the Northern Hemisphere. However, certain species have been identified in both polar regions and others in warmer locales like Queensland (Australia). Schaereria species often grow on siliceous rocks, with a few variants being epiphytic (plant-dwelling) or even lichen-dwelling. Various lichenicolous fungi and lichens have been observed to grow on Schaereria species.
Genus Schaereria was named and circumscribed by the German lichenologist Gustav Wilhelm Körber in 1855, with Schaereria lugubris assigned as the type, and at the time, only species. [3] According to Hannes Hertel and L. Zürn, it is clear that he was actually referring to Schaereria cinereorufa(Schaer.) Th.Fr., but he mistakenly believed he had Ropalospora lugubris (formerly Lecidea lugubris) on hand. For this reason, Körber's name is not available for the genus. [4] Despite this, Schaereria was accepted a few years later (in 1860) by Theodor Magnus Fries, who used it in his work on Arctic, European, and Greenlandic lichens. [5] It subsequently fell into disuse for more than a century after William Nylander placed it in synonymy with Lecidea . [6] Josef Poelt and Antonín Vězda resurrected Schaereria in 1977, and included S. cinereorufa . [7] The genus name honours the Swiss pastor and lichenologist Ludwig Emanuel Schaerer. [8]
In 1989, David Leslie Hawksworth and John Charles David put forth a proposal to conserve the name Schaereria (as defined by Theodor Magnus Fries in 1874) for a group of species with S. cinereorufa as the type species, instead of using Schaereria as defined by Körber. The reason for this proposal is that S. cinereorufa is a well-known species under the name Schaereria, while Körber applied the name S. lugubris to a different species that should have been named S. cinereorufa. By conserving the name Schaereria as suggested, it avoids the need to create a new name for the same group of lichens and prevents confusion with another genus called Ropalospora. By accepting this proposal, both Ropalospora and Schaereria could continue to be used with their current meanings, avoiding any unnecessary changes. [9] In the subsequent recommendations of the Committee for Fungi and Lichens, they agreed that it was desirable to conserve the genus with one of the original specimens used by the author, noting that this specimen was erroneously labelled Schaereria lugubris, but clearly belongs to S. cinereorufa. [10]
Schaereria is one of several dozen genera whose species were previously included in the large genus Lecidea. [11] However, Lecidea has a different ascus structure than Schaereria. The family Schaereriaceae was first proposed by French lichenologist Maurice Choisy in 1949, but he did not publish the name validly as it did not meet the criteria for publication as determined by the International Code of Nomenclature for algae, fungi, and plants. [12] Josef Hafellner published the family name Schaereriaceae validly in 1984. [13]
Hafellner noted some similarities in the characteristics of the hymenium between the Schaereriaceae and the order Pezizales, [13] and the family was included there in the 1985 version of the Outline of the Ascomycota. [14] This classification was later shown to be inappropriate as the Schaereriaceae do not have operculate asci—a characteristic of the Pezizales. After this the family was associated with either the Teloschistineae (a suborder of the Teloschistales), the Agyriineae (a suborder of the Lecanorales), [6] or placed in the order Sarrameanales. [15] Schaereria has also been included and excluded from the family Lecideaceae several times. [16] Molecular studies that have included Schaereria species have shown that it occupies a relatively isolated phylogenetic position. [17] [16] [18]
In 2018, H. Thorsten Lumbsch and Steven Leavitt proposed the new order Schaereriales to contain the family, segregating the order Schaereriales from Sarrameanales. They considered the Schaereriales to have "unresolved relationships" in the parent taxon, Ostropomycetidae. In their analysis, which used a "temporal" method to group together comparable ordinal and family ranks, Schaereriales has a sister group relationship with the order Sarrameanales. Both Schaereriales and Sarrameanales form a clade that is sister to the Baeomycetales. [19] [20] In a critical review of the temporal method for lichen classification, Robert Lücking found flaws in their analysis and rejected the proposed split, instead retaining both Sarrameanaceae and Schaereriaceae in the Sarrameanales. [21]
Several characteristics unite taxa in the order Schaereriales. These include: a thallus that is crustose to squamulose; a trebouxioid photobiont partner (spherical unicellular green algae); ascomata being lecideine (having an apothecium which lacks algae and lacks an amphithecium ); apothecia that are hemiangiocarpous (meaning they open before the spores are mature); and a cup-shaped excipulum . Microscopic characteristics of the Schaereriales include asci of the Schaereria-type (lacking a tholus –the thickened inner part of an ascus tip), and ascospores that are hyaline, thin-walled, and one-celled (i.e., lacking septa). [19] There are eight spores per ascus. The hamathecium contains non-amyloid, non-gelatinous, unbranched paraphyses . [15] The secondary chemistry of this group includes depsides and unknown compounds. [15] Schaereria cinereorufa produces a green- to turquoise-coloured insoluble lichen pigment known as Cinereorufa-green . [22] The identification of species within Schaereria is primarily based on specific traits: the chemical composition and form of the thallus, the colouration of the apothecia, and the distinctive shape and organization of ascospores within the ascus. [23]
The anatomy and development of several Schaereria species have been studied in some detail. The genus displays a varied morphology, encompassing both crustose and squamulose lichens. For instance, Schaereria cinereorufa is characterised by a squamulose to bullate (blistered) thallus, often found on siliceous rocks, with sessile or marginally located apothecia. In contrast, S. corticola has a sparsely developed, endo- to episubstral thallus with soredia and sessile apothecia. Schaereria fuscocinerea, somewhat intermediate, features an areolate thallus on siliceous rocks, and its fruiting bodies are either immersed in or between areolae . [6]
The anatomy within the genus is more consistent. In S. cinereorufa and S. fuscocinerea, the thallus is covered by an epinecral layer (a layer of dead fungal hyphae), and no distinct cortex is formed. However, the upper parts of the hyphae in the thallus are pigmented brownish. In contrast, S. corticola lacks a distinct upper layer, primarily consisting of soredia. [6]
The development of Schaereria ascomata is somewhat variable depending on species. In S. cinereorufa, the ascomatal primordium begins as groups of ascogonia forming a more or less spherical cluster, followed by coiled ascogonia with trichogynes . The primordium eventually rises to the thallus's upper surface, with the upper part forming the hymenium. In S. corticola, spherical primordia with ascogonia are observed, becoming pigmented and forming the cup-shaped excipulum . True paraphyses replace the paraphysoidal network in the hamathecium . In S. fuscocinerea, ascogonia with trichogynes appear in the algal layer , with subsequent growth of paraphysoids, followed by the differentiation of ascogenous hyphae and true paraphyses. [6]
Collectively, the genus has a cosmopolitan distribution. [15] The majority of Schaereria species inhabit cold to cool climates within the Northern Hemisphere. S. fuscocinerea, however, is an exception, as it can be found in both polar regions. Additionally, S. xerophila thrives in the lowland regions of Queensland. Schaereria species are typically found growing on rocky surfaces composed of siliceous materials. In contrast, S. corticola is bark-dwelling, S. dolodes is both bark- and wood-dwelling, [16] S. corticola has an epiphytic lifestyle, while S. parasemella has a lichenicolous (lichen-dwelling) habit. [24]
Some lichenicolous fungi and lichens have been recorded growing on Schaereria species. These include Endococcus perpusillus , Buellia miriquidica , Buellia uberior , and Halecania parasitica (the last three on Schaereria fuscocinerea). [25] A Sclerococcum species in the S. australe – S. saxatile group has been reported infesting Schaereria bullata. [24]
As of January 2024 [update] , Species Fungorum (in the Catalogue of Life) accepts 11 species of Schaereria. [26]
Several taxa once classified in Schaereria have since been reduced to synonymy, or transferred to other genera. Examples include:
The Lecanoraceae are a family of lichenized fungi in the order Lecanorales. Species of this family have a widespread distribution.
Gloeoheppiaceae is a family of ascomycete fungi in the order Lichinales. The family contains ten species distributed amongst three genera. Most species are lichenised with cyanobacteria. Species in this family are mostly found in desert areas. Modern molecular phylogenetics analysis casts doubt on the phylogenetic validity of the family, suggesting a more appropriate placement of its species in the family Lichinaceae.
The Baeomycetales are an order of mostly lichen-forming fungi in the subclass Ostropomycetidae, in the class Lecanoromycetes. It contains 8 families, 33 genera and about 170 species. As a result of molecular phylogenetics research published in the late 2010s, several orders were folded into the Baeomycetales, resulting in a substantial increase in the number of taxa.
Stirtoniella is a lichen genus in the family Ramalinaceae. It is a monotypic genus, containing the single species Stirtoniella kelica, a crustose and corticolous lichen originally described from New Zealand in 1873 as a species of Lecidea. The photobiont is an alga of the family Chlorococcaceae. The genus is named after Scottish mycologist James Stirton.
Malcolmiella is a genus of lichenized fungi in the family Pilocarpaceae.
Lecidella is a genus of crustose lichens in the family Lecanoraceae.
Ramboldia is a genus of lichen-forming fungi in the family Ramboldiaceae. The genus was circumscribed in 1994 by Gintaras Kantvilas and John Alan Elix. It was emended in 2008 by the inclusion of Pyrrhospora species containing the anthraquinone russulone in their apothecia and having a prosoplectenchymatous exciple. The family Ramboldiaceae was circumscribed in 2014 to contain the genus.
Porpidia is a genus of crustose lichens in the family Lecideaceae.
Rhizocarpon is a genus of crustose, saxicolous, lecideoid lichens in the family Rhizocarpaceae. The genus is common in arctic-alpine environments, but also occurs throughout temperate, subtropical, and even tropical regions. They are commonly known as map lichens because of the prothallus forming border-like bands between colonies in some species, like the common map lichen.
Fuscidea is a genus of crustose lichens in the family Fuscideaceae. It has about 40 species. The genus was circumscribed in 1972 by lichenologists Volkmar Wirth and Antonín Vězda, with Fuscidea aggregatilis assigned as the type species.
Hypocenomyce is a genus of lichen-forming fungi in the family Ophioparmaceae. Species in the genus grow on bark and on wood, especially on burned tree stumps and trunks in coniferous forest. Hypocenomyce lichens are widely distributed in the northern hemisphere.
Malmideaceae is a family of crustose and corticolous lichens in the order Lecanorales. It contains eight genera and about 70 species.
Varicellaria is a genus of crustose lichens. It is the only genus in the family Varicellariaceae.
Rhizocarpales are an order of lichen-forming fungi in the subclass Lecanoromycetidae of the class Lecanoromycetes. It has two families, Rhizocarpaceae and Sporastatiaceae, which contain mostly crustose lichens.
Strangospora is a genus of lichen-forming fungi. It is the only genus in the family Strangosporaceae, which itself is of uncertain taxonomic placement in the Ascomycota. It contains 10 species.
Schaereria fuscocinerea is a species of saxicolous (rock-dwelling), crustose lichen in the family Schaereriaceae. It was first formally described in 1852 by Finnish lichenologist William Nylander, as Lecidea fusco-cinerea. Georges Clauzade and Claude Roux transferred it to the genus Schaereria in 1985. The species has a cosmopolitan distribution and is found in both northern and southern hemispheres, where it grows on hard siliceous rocks, often in arctic and mountainous areas. Similar species include Lambiella gyrizans and L. mullensis, which can be distinguished from Schaereria fuscocinerea by microscopic and chemical characteristics.
Schaereria serenior is a species of saxicolous (bark-dwelling), crustose lichen in the family Schaereriaceae. Found in Finland, it was first formally described as a new species by the Finnish lichenologist Edvard August Vainio, who classified it as a variety of the species Lecidea tenebrosa. Auguste-Marie Hue promoted it to distinct species status in 1913 as Lecidea serenior. Alexander Zahlbruckner proposed to transfer it to the genus Caloplaca in 1931. Most recently, Orvo Vitikainen transferred it to Schaereria in 2004, a few years after that genus had been resurrected from a long period of disuse. It is one of two species of Schaereria found in Finland; the other is S. parasemella.
Schaereria parasemella is a species of saxicolous (rock-dwelling), crustose lichen in the family Schaereriaceae. It is lichenicolous, meaning it grows on other lichens; its host species is Biatora vernalis. It has been recorded from Alaska, Greenland, Scandinavia, France, and Russia.